Botanica Marina Vol. 45, 2002, pp.


© 2002 by Walter de Gruyter · Berlin · New York

A Revision of Amansia glomerata C. Agardh, Amansia rhodantha (Harvey) J. Agardh and Melanamansia glomerata (C. Agardh) R. E. Norris (Rhodophyta: Rhodomelaceae)
A. D. R. N’Yeurt
Jeune Equipe Terre Océan, Université de Polynésie Française, BP 6570, Faa’a Aéroport, Tahiti 98702, French Polynesia,

Species of Amansia and Melanamansia from the tropical Pacific region have been re-examined, and compared with material from the Indian Ocean, Japan, Taiwan and the Philippines. All specimens examined from the Pacific region (except Hawaii and some from New Caledonia and Fiji) lack pseudo-pericentral cells, thus placing them in Amansia. Melanamansia glomerata is only reported from Hawaii, New Caledonia, Fiji, the Philippines,Vietnam, Malaysia and for the first time in the Indian Ocean from Kenya.Type material of Amansia rhodantha (Harvey) J. Agardh from Mauritius was compared and found not significantly different from the Pacific Amansia material (including type and isotype material of Amansia paloloensis South et Skelton from Samoa, which is reduced to synonymy), thus extending the distribution range of A. rhodantha as a pantropical entity.

The genus Amansia was erected by Lamouroux (1809: 322) and is placed in the tribe Amansieae of the family Rhodomelaceae (Hommersand 1963: 335). The tribe is characterised by dorsi-ventral, erect thalli arising in tufts from a common basal disc. Most genera have five pericentral cells around the axial cell (Figs 1, 2) except Kuetzingia Sonder and Protokuetzingia Falkenberg, which have six pericentral cells (Kylin 1956). The two first-formed pericentral cells lie on the dorsal side of the characteristically inrolled apex of the mature lanceolate blades. In Amansia the dorsal and dorsi-lateral pericentral cells further divide to form immediately behind the apex a distromatic wing or ala, whose cells are arranged in alternate layers, with secondary lateral pit-connections (Fritsch 1945: 570). Recently, Norris (1988b, 1995) re-examined species of Amansia in the Indian Ocean and Hawaii, and discovered that some members of this genus differed consistently by having the first and second pericentral cells each producing a lateral pseudo-pericentral cell lying adjacent, but not connected to, the axial cell (Norris 1988b: 214; Fig. 3). Species previously ascribed to Amansia that had these pseudo-pericentral cells were transferred to a new genus (Melanamansia R. E. Norris 1988b: 217). Notably, Hawaiian plants previously ascribed to the common tropical species Amansia glomerata C. Agardh (whose type locality is Hawaii) were transferred to Melanamansia glomerata (C. Agardh) R. E. Norris, while Indian Ocean ‘Amansia glomerata’ were put under the earlier name Amansia rhodantha (Harvey) J. Agardh based

on their lack of pseudo-pericentral cells (Norris 1995). The genus Amansia currently consists of four species, the Caribbean A. multifida Lamouroux (the type of the generic name), A. loriformis R. E. Norris and A. rhodantha (Harvey) J. Agardh from the Indian Ocean and Malaysia, and A. paloloensis South et Skelton from Samoa. Melanamansia currently consists of twelve species, M. daemelii (Sonder) R. E. Norris, M. glomerata (C. Agardh) R. E. Norris, M. japonica (Holmes) R. E. Norris, M. mamillaris (Lamouroux ex C. Agardh) R. E. Norris, M. pinnatifida (Harvey) R. E. Norris, M. pumila (Sonder) R. E. Norris, M. scalpellata (Tanaka) R. E. Norris, M. seagriefii R. E. Norris, M. serrata (Harvey) R. E. Norris and M. mitsuii (Tanaka) Yoshida. Following these revisions by R.E.Norris,several authors (Yoshida et al. 1995,N’Yeurt et al. 1996,Payri and N’Yeurt 1997, Millar 1999, Millar et al. 1999) made the nomenclatural change from Amansia glomerata to Melanamansia glomerata in their checklists and herbarium collections from the tropical Pacific, but until recently no detailed studies verifying these changes as correct have been done. In 1995, samples of ‘Amansia glomerata’ from Rotuma Island (north of Fiji) were sent to Dr R.E.Norris,who confirmed (pers. com.) that pseudo-pericentral cells were present, and on the basis of this assumed finding the name Melanamansia glomerata was used in later publications (N’Yeurt 1996,N’Yeurt et al. 1996).Later,in the course of on-going investigations on the Samoan algal flora, South and Skelton (1999) found a species of Amansia lacking pseudo-pericentral cells, which they described as new (A. paloloensis South et Skelton), while in Malaysia and Vietnam (Masuda et al. 2000, Masuda

com. A. 2 Jul.) recent studies found both Melanamansia glomerata and Amansia rhodantha (Harvey) J. leg. 11 Jul. 3899). Afiti and D. Baba (SAP 056160). a detailed morphological and taxonomic study on some selected members of the Amansieae (Wilson and Kraft 2000) echoed the opinion that the genus Amansia in Australia was in need of critical revision. Amansia rhodantha and Amansia glomerata. a = axial cell. leg. cross section of the lectotype from the Hawaiian Islands (LD 42600) showing the first two dorsal pericentral cells each developing a pseudo-pericentral cell (ps) that lies adjacent to the axial cell. snorkelling or SCUBA diving. A. – Taiwan: Anon. leg. 1998. N’Yeurt (SUVA-A 5128). Fig. 3892. P. leg. Skelton. 1984. A. Arrangement of pericentral cells in species of Amansia and Melanamansia (modified from Norris 1988b. Tombleson and T. 1998. Fiji. 6 Jun. showing intrusion of the first derivative of the first-formed pericentral cell into the axial ring (Suva. Melanamansia glomerata. Most recently. 13 May 1982. and stained with cotton blue in lactophenol or 1% acidified aniline blue in 60% clear corn syrup. 25 October 1984 (SAP 046978). Skelton. P. leg. 1. SUVA-A 5438L). SUVA-A 5436L). 3898. Okinoerabujima Island (60 km north of Okinawa) 27 Sept. Amansia rhodantha (Beqa. N’Yeurt (UPF 347). Rintels (SUVA-A 3891. otherwise as in Holmgren et al. Tahiti. Skelton and Dave (SUVA-A 1567). . A. leg. E.‘Al’ and ‘CG’ refer to specimens housed at IRD. N’Yeurt In order to clarify the taxonomy of species hitherto ascribed to Melanamansia glomerata. Fiji. leg. Skelton and Dave [SUVA-A 1565 (holotype). 1998. – Samoa: Palolo Deep Reserve. U. Palolo Deep. Macrophotography was done with a Nikon F2A in conjunction with Kodak Plus-X pan 125 film. A. A. SUVA-A 2402 (Lectotype) and liquid-preserved isotypes 5479L (tetrasporic). 18 Sep. 5481L (spermatangial)]. and a camera-lucida attachment was used to make anatomical drawings. 1989. D. M. M. P. Taharaa. ps = pseudo-pericentral cells). Figs 1–3.. Photomicrographs were obtained using an Olympus BX-50 compound microscope. it was decided to revise records in the light of recent findings. 3896. 1999. and stored in 5% buffered Formalin in seawater prior to shipment to the laboratory. Okidomari. deep algal rim. 1997. A. M. 1–5 = order of formation of pericentral cells.232 A. The results of these investigations are described below. leg. C. 9 Feb. Payri (8/84. Enomoto (SAP). Furusato. Sakai and S. Y. 1998. 1998. SUVA-A 1566 (isotype). developed and printed in the laboratory. P. Tanegashima Island. in EPHE). Faletose (SUVA-A 3894. in particular paying attention to important anatomical features such as the presence or absence of pseudo-pericentral cells and the number of segments between lateral veins of the blade (Norris 1995). Okinoerabujima Island (60 km north of Okinawa). – French Polynesia: Moorea. 3. Amansia rhodantha. Rarotonga. Prepared slides were examined using a Zeiss compound microscope. 19 Mar.Agardh to co-exist in the flora. 2 Jul. pers. Punaauia. leg. R. leg. N’Yeurt (UPF 584). 3897). Nouméa. Sections were made using a freezing microtome. New Caledonia. Material and Methods Specimens were collected by reef-walking. 3895. Kagoshima: 9 August 1959. – Cook Islands: Ngatangiia. leg. 2. 3893. Fig. Tahiti. Results Representative material examined Amansia rhodantha – Japan: Hanasado Beach. Baba (SAP 056163). Fig. (1981). 5480L (cystocarpic).

M. Species Thallus height (cm) Blade width (mm) Blade length (mm) Serrations (1 –3)* Cortication (1 –3)* Rosettes (1 –3)* Amansia rhodantha Amansia paloloensis Melanamansia glomerata 39. A. leg. 84 (L 0108603). Cribb (SAP 051714). leg. Oahu. R. 1876. Skelton and G. Cannon (SAP 052062). Hakula. Gaudichaud (‘Amansia glomerata’. Kipukai. leg. leg. paloloensis and Melanamansia glomerata specimens examined in this study. Richer de Forges (AL 288d-CG 77a). Huisman (MURU. R. – Fiji: ‘Feejee’. Average values of selected characters for Amansia paloloensis. Yasawa Islands.. 10 Aug. Snellius-II (L 0108602). leg. Banilad. leg. M de Robillard (L 0108599). Oahu. leg. 15 Oct. 1995. 3 Jun 1978. leg. M.P. G. L 0108598). 17 Mar. 5 June 1967. Agardh 1841: 26.Pagodpud. Tantabiddi. 14 Feb. leg. Farlow (L 0108597). Huisman (MURU. C.. Ovalau. Sumbawa. Inhaca Peninsula. leg. N’Yeurt (SUVA-A 346. leg. – Mauritius: Delesseria rhodantha Harvey (holotype) (= Amansia rhodantha): Cap Malheureux. leg. Lawai Kai. G. 14 March 1998. 17 Jul. B.. Huisman (CWC 1997. leg. 10 July 1992. leg. – New Caledonia: Lajeu Sudouest. B. J. – Lord Howe Island: 18 Aug. 1984. Viti Levu. Keats. Viti Levu. 1838–42. 07 Oct. Hanauma Bay Beach Park. 1067). 1819. 25 September 1999. – Indonesia: Cape Batu Kerapo. W. W.. leg. Silva et al. J. Suva Reef. 3 and 15 Oct. SIPHILEXP-78 (US 088531). (MURU. Norris 1988b: 211. Garrigue (CG 77c.14 May 1979. 1988. D. A. – Fiji: ‘Feejee’. 1979. Tangangge. 84. – Mozambique: Ponta Abril.Palawan. n. G. M. Kauai. W. S. Negros Oriental. Vatualailai. Nanuyalevu.7 2 2 1 2 2 2 2 2 2 * As measured on an index of (1 = little) to (3 = pronounced) – Rotuma: Hoféa. n.. Cordero and Tuangpalan (US 091394). Beqa Lagoon. Villeneuve (SUVA-A 1774). B79). Jul. leg. SIPHILEXP-79 (US 088533). P. leg. M. 25 Feb. 18 March 1992. R. Wilkes Exploring Expedition (US 04057b) (doubtful record.Negros Oriental. figs 37–42. leg. C.d.. in TCD (Telfair. – New Caledonia: Lajeu Sudouest. 16 Nov. B.Revision of some Amansia and Melanamansia species 233 Table I.Calindagan Reef. Yasawa Group. CG 77b). 1985. – Australia: Heron Island QLD. Barrow Island. R127). 1995. 1995. 1981. leg. B262). 347). Magnesia Virae. Carlson (SUVA-A 281).W. Daida (US 086755). leg. SIPHILEXP-78 (US 013760). n. A. leg. 2000: 188.9 18. A. leg.. anon. leg. c. Huisman. C. 84. Isaac (L 0108595). 1838–42. 1965. leg.d. H. LD 42600 / 0772). 1834). 1982. leg. 1997. Bahati. leg.leg.Solong-On. leg. leg. N’Yeurt (SUVA-A 5437L). leg. 21 Sep. 26 Nov. South (SUVA-A 5544. J. Unia. Cordero and Tuangpalan (US 091395).Canipo Is. Negros Oriental. Snellius-II (L 0108601). 1876. Montebello Islands.23 May 1978. 3 and 5 March 1992.. R.Malo. Potipot Is.6 3. Masuda et al. A. Namada. Basay. leg. 1983. Mauritius. 1994 (SUVA-A 797). Melanamansia glomerata – Hawaiian Islands: Hawaiian Islands. leg. Agardh). 10 May 1978. 6 Jun 1978. J.11 Feb. Isaac (L 0108594). Amansia rhodantha and Melanamansia glomerata surveyed in this study. JH240). Hapmak. 1979. leg.leg. A. A. Hildebrandt (L 0108596.3 2. S. 9 Sept. leg. South (SUVA-A 5431). Lectotype. Naukathura Island. 1984. 1069).2 53. Chesterfield. Ballou (SUVA-A 5438L). Viti Levu.Dumaguete City. – Philippines:Gao-oa.Siquijor Is. H. 30 Jun. Garrigue (SUVAA 5436L). Kauai. W. n. Garrigue (CH1). 24 April 1996. Oahu.9 11. Bay of Sanggar. 3 Feb. leg. Table I presents the average results of morphological measurements carried out on Amansia rhodantha.1 22. Barrow Island. 16 Oct. C. SIPHILEXP-78 (US 088532). 1973.). Figs 4–10 .d. Magruder (US 086459).5 3. Ti Tree.Ilocos Norte. E. figs 1–11 (as Amansia glomerata C. J. Candelaria Zambales. A. Suva Barrier Reef. leg. 13 July 1972.Negros Oriental. (US 02890). Salayer. H.7 54. 16 March 1991. Hildebrandt (L 0108600). leg. 1985. leg. 1980. fide G. leg. E. Taxonomic results Amansia rhodantha (Harvey) J. Littler (US 163490). J. Kasahara (SUVA-A 1012.Siaton. R. P.d. SIPHILEXP-78 (US 088534). leg. NR 248). 24 Nov. Ningaloo. 21/22-Sept. Catanduanes. leg. Rukuruku Village. E.d. Wilkes Exploring Expedition (US 04057a). Huisman (MURU. South in lit. SIPHILEXP-78 (US 088530). Jurien Bay WA. G. South (SUVA-A 2460). leg. Jul. 1954. Villeneuve (SUVA-A 1737). 1996: 472. 06 Sept. 29 May 1978. Dumaguete. leg. P. 27 Dec. Diani Beach. 1982.Moreland (US 50851). – Kenya: Mombassa. Lopta. – Philippines: Actin. leg. 28 Nov.Cuyo. Kasahara (SUVA-A 1013. Amansia rhodantha: n. 1980. leg. J. leg. see text). – Kenya: Mombassa. AL 288b). Eagles Nest. leg. Plongeurs IRD (AL 288a.

SUVA-A 5437L. 10. Axial structure of blade. in TCD (scale bar = 200 µm). R. . in TCD (scale bar = 25 µm). Secondary branchlet. D. South Pacific (scale bar = 10 µm). Mauritius. showing marginal serrations.234 A. Type material of Delesseria rhodantha Harvey. Indian Ocean. Surface view of segment cells of blade alae. 8. N’Yeurt Figs 4–10. Figs 4. 5. Rehydrated type material of Delesseria rhodantha Harvey. from Rotuma Island. in TCD (scale bar = 2. Fig. Fig. Habit of mature plant from the type locality. 7. Fig. Detail of tetrasporangial stichidia on lateral endogenous branch. Amansia rhodantha. US 02890 (scale bars = 3 mm). 9.5 mm). Fig. Type material of Delesseria rhodantha Harvey. showing axial cell (a) surrounded by five pericentral cells. showing both lax and dense growth forms. and a pair of budding branchlets from the midrib in the lower half. 6.

15. showing lateral endogenous branches and midrib. SUVA-A 3892 (scale bar = 2. Fig. SUVA-A 5436L (scale bar = 50 µm). SUVA-A 5479L (scale bar = 1 mm). Detail of morphologically distinct mature rosette and corticated axis. 16. SUVA-A 3892 (scale bar = 12 mm). Fig. 13. Fig. 11. Holotype of Amansia paloloensis. SUVA-A 1565 (scale bar = 25 µm). Fig. Fig.5 mm). Compare with Fig. Fig.Revision of some Amansia and Melanamansia species 235 Figs 11–16. 12. showing both lax branching and the presence of rosettes (arrowheads) (scale bar = 4 mm). Amansia paloloensis from Samoa. Note marginal cystocarps (arrowheads). 4. Detail of blades. Detail of the holotype (SUVA-A 1565). with numerous rosette axes. Tetrasporangial stichidia of Amansia rhodantha from Beqa. . Habit of mature plant. Segment cells of blade alae in surface view. Fiji. 14.

composed of two overlapping layers of elongate hexagonal cells in transverse rows. US 091395 (scale bar = 5 mm). pl.4. The thallus is rose-red. L 0108598 (scale bar = 5 mm). Branching is lax to profuse and usually forms deep-red leafy rosettes of secondary branchlets when fully mature. Falkenberg 1901: 416. 18. 19. Carposporangia are lanceolate. Fig.8 mm in diameter. and are composed of lanceolate to ovate blades to 100 mm long and 3 mm wide. 85–110 µm thick. Figs 17–24 Basionym: Amansia glomerata C. Tropical Pacific Ocean. Philippines. Vietnam. e–i. Kenya. Synonymy: Rytiphlaea rhodantha (Harvey) Decaisne 1842: 358. Cribb 1983: 106. Payri and N’Yeurt 1997: 897. Fig. Distribution: Tropical Indian Ocean. Kenya. . Fig. 20. The main axis is irregularly branched. 267. Philippines. 31 fig. Japan. with a range of 3–6. A distinct midrib up to 1 mm wide is present in the lower half of ultimate branchlets. Plant from Potipot Island. Norris 1995: 67. Procarps and curved spermatangial heads are 200–450 µm in diameter and are dorsal on the involucral tips of endogenous lateral branches. The central axial cell is 28–30 µm in diameter. holotype in TCD. pinnately arranged marginal stichidia up to 340 µm long and 290 µm in diameter. East Africa. Blades of plant assumed to be from the Fiji Islands (see text). Fig. Individual tetrasporangia are spherical and tetrahedrally divided. 212a. 23. AL288b (scale bar = 5 mm). Trichoblasts are sparse.5 µm in diameter that lies adjacent to the axial cell (Figs 3. – Melanamansia glomerata (C. and are crisp and lanceolate. E. Tetrasporangial stichidia are pinnately arranged on either side of blade margins. Tetrasporangial stichidia of plant from Oahu. Okamura 1900: 71. Dublin). 118C–J. becoming narrower and disappearing towards the apex. pl. Figs 11–16 Misapplied names: Amansia glomerata C. New Caledonia. Garrigue and Tsuda 1988: 63. figs 1–3 (type locality: Cap Malheureux. 100–112 µm in diameter. and are initially vesiculate. 6E (unverified record). Jaasund 1976: 131. 198. 17. with more or less pronounced marginal serrations 1–4 mm long (which represent endogenous branches) always present. Deciduous. A central midrib is present. Abbott 1999: 404. plate CXXVI. 1996: 87. 24. Known distribution: Hawaiian Islands. Ultimate branchlets are ecorticate. E. figs 1–13. 9). in up to 10 regularly disposed pairs. New Caledonia. Fig. 1. Apia. Plant from Unia. with in-rolled leaf apex. colourless and uniseriate dorsal trichoblasts up to 8 cells long develop on every second or third axial cell (or on every segment in determinate marginal branchlets). Fiji. 77–110 µm thick. fig.5–0. Norris 1995: 66) is on average 4. showing pseudo-pericentral cells (arrowheads) surrounding (but not directly connected to) axial cell (a). 4B. The stem of plants is thick and cartilaginous. denuded below. up to 100 (on average 40) mm high. Melanamansia glomerata (C. with a central axial cell 28–30 µm in diameter surrounded by five pericentral cells 36–40 µm in diameter (Figs 1. to 0. containing up to 10 Figs 17–24. XXV. on average 53 mm high. pl. 1999: 573. Fig. figs 127. and the apex is characteristically in-rolled. showing smooth margins and elongate shape. Yoshida et al. 22. Norris 1995: 66) is usually 4. Payri and Meinesz 1985: 511. Coppejans and Millar 2000: 333 (unverified record). New Caledonia. figs 19–21. Internal structure is cellular.8–14. Blade of plant from Lajeu. 1987: 61. 0. Isaac 1956: 188. N’Yeurt 1996: 428. 7 mm broad. Detail of blade of plant from Mombassa. Agardh) R. Yamada and Tanaka 1938: 86. or with regular serrations (endogenous branches). fig. Plants are brownish-red. Samoa). CG77a (scale bar = 1 mm). N’Yeurt et al. 19). 25–42 × 83–125 µm. Millar 1999: 523 (unverified record). South-East Asia. Fig. 20. D. stem-like and rigid in basal portions. pl. up to 750 µm long and 250 µm broad. Axial structure of plant from Lajeu. while cystocarps 700–1000 µm in diameter occur usually singly and terminally on endogenous branches. CG77a (scale bar = 10 µm). 197. fig. figs 1–11. Kylin 1956: 544. 128. they are up to 4 cells long and non-vesiculate. figs 14–29. each cell is about 20–20 × 87–100 µm in surface view. N’Yeurt Basionym: Delesseria rhodantha Harvey 1834: 151–152. 436B. fig. Marginal teeth are up to 1 mm long and 445 mm broad. Fritsch 1945: 570. Note that the stichidia is twice laterally branched. fig. pl. with a range of 1–5. Secondary blades develop as successive pairs on the dorsal side of the midrib. R. figs 2–14 (type locality: Palolo Deep Marine Reserve. 1. 21. with the first two dorsal pericentral cells each developing a smaller pseudo-pericentral cell 13. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. Agardh 1824: 194. Norris 1995: 67.236 A. fig. Melanamansia glomerata. The number of segments between alternating veins (determinate branches covered by the corticating pericentral cell derivatives. Agardh 1824: 247. SUVA-A 5431 (scale bar = 50 µm). US 4057b (scale bar = 1 mm). Mauritius. Millar et al. Agardh) R. Tsuda and Wray 1977: 103. New Caledonia. up to 35 mm long and 6 mm broad. Tetrasporangia occur in flattened. and two displaced rows of regularly arranged hexagonal alar cells 20–29 × 92–129 µm in surface view. Womersley and Bailey 1970: 336. Isaac and Isaac 1968: 24. L 0108598 (scale bar = 700 µm). 6. surrounded by five pericentral cells. Lewis and Norris 1987: 22. 1995: 150. 185. 2. Kenya. Silva et al. Norris 1988b: 211. Fig. Amansia paloloensis South et Skelton 1999: 247. Plant from Mombassa. Blade margins are smooth. Weber-van Bosse 1923: 369.

Revision of some Amansia and Melanamansia species 237 pairs of tetrahedrally-divided tetrasporangia which are 50–100 µm in diameter. procarps and cystocarps occur on endogenous marginal branches. . and stained the herbarium mounting paper. the colour of the Melanamansia plants was noticeably brownish-red. Spermatangial heads are curved and arranged in marginal clusters. In all cases.

a re-examination of the lectotype of Amansia glomerata C. a single record for this genus exists (collected from ‘Feejee’ by the Charles Wilkes U. isotype and other collections of A. Now that the Samoan species has been compared with genuine herbarium material of A. and Ceramium. loriform blades reminiscent of some specimens of A. N’Yeurt Discussion The holotype of Delesseria rhodantha Harvey An examination by the author of a rehydrated blade fragment of the holotype of Delesseria rhodantha Harvey in TCD (Fig. Vidalia. paloloensis. Kenya. Rotuma. M. having tight rosettes as well as very lax. MURU 248). and a cross section of the blade clearly shows the five pericentral cells around the axial ring. Agardh from Hawaii (LD 42600 / 0772) clearly shows the presence of paired dorsal pseudo-pericentral cells in a blade cross section (Fig. and absence of pseudopericentral cells.13). The present author has re-examined the holotype. Mozambique and Western Australia. Smith (1979: 38–40) points out that the accounting and . it appears to be conspecific with the latter species and thus extends the status of A. a comparison of material of A. 3) which distinguishes that plant from A. However. rhodantha as described and illustrated by Norris (1988b: 211). In the Fiji Islands. including the presence of both tight and lax rosettes of secondary branches and alar cells on the primary axes. Fiji. fig. and these authors considered the lack of highly corticated axes. Masuda pers. The Wilkes collection consists of two plants. material from the Tropical Pacific and Indian Oceans previously known as Amansia glomerata (or misidentified as Melanamansia glomerata. paloloensis.238 A. additional distinguishing characteristics of Melanamansia are a darkly coloured brown thallus which stains the mounting paper when dried (possibly linked to differences in chemical composition. regularly alternate marginal serrations which are not as pronounced as in Amansia rhodantha and which are often absent. from Melanamansia glomerata was found: when preserved in 4% Formalin-seawater and exposed to ambient light for several days. The reproductive and trichoblast anatomy of A. rhodantha from the Indian Ocean and other localities.S. the Philippines. These plants also had blades with well-developed midribs in the lower half (Fig. plate CXXVI. From this study. D. is comparable to A. The drawing of the holotype (South and Skelton 1999: 246. serrate branches and lax ‘rosettes’ of secondary blades issued from the weakly developed midrib. the absence of blades organized in morphologically distinct tight axes around the parent axis. 23) but the plant has not been collected in Fiji waters since. while Melanamansia becomes dark brown. figs 1–4) indicate features quite similar to those found in the Amansia species considered in this study from localities in both the Indian Ocean and the tropical Pacific Ocean. and a weakly developed midrib as characters distinguishing A. rhodantha (South and Skelton 1999: 249). rhodantha from the type collection and localities in Mauritius. Kuetzingia. which is welldocumented and illustrated in the protologue of the species. While there is no doubt as to the generic identity of the latter specimen. 7) shows features similar to those seen in blades of South Pacific material previously ascribed to Amansia glomerata. The habit and description of the holotype by Harvey (1834: 151. The status of Amansia paloloensis South et Skelton South and Skelton (1999) described a new species of Amansia from Samoa. generally smaller. degree of cortication of the midrib and position of the reproductive stichidia are equally quite variable in plants from the same and similar populations. when pseudo-pericentral cells were found to be absent) are considered conspecific with Amansia rhodantha. com. Exploring Expedition. Characters such as the degree of rosette formation. Melanamansia glomerata in the tropical Pacific Wilson and Kraft (2000: 366) commented that the generic segregation of Melanamansia from Amansia is probably not warranted. paloloensis from A. length of the blades. 2) indeed shows a plant with loriform. R. rhodantha. between 1838 and 1842 (US 04057b. On the basis of these observations. with for instance A. New Caledonia. paloloensis showed no major difference in the range of characteristics among them. since the presence of pseudo-pericentral cells is an inconsistent attribute in species of Neurymenia. rhodantha from Western Australia (MURU B262. 14) and corticated rosette axes (Fig. Fig. More importantly. rhodantha as a pan-tropical entity. 12). and found a significant number of thalli with morphologically distinct primary corticated axes and relatively tight and lax rosettes of secondary blades on rosette axes (Figs 11. one of which (US 04057a) is Amansia rhodantha. as well as Amansia species from Taiwan. paloloensis from the type locality. apparently because the brown pigments (presumably polyphenols) are less susceptible to the bleaching action of the preservative (as is the case in species of brown algae such as Sargassum and Turbinaria). Eastern Australia.). indicating that the morphology of the species is influenced perhaps by environmental factors. more coriaceous branchlets with less apparent midribs (this study). the Cook Islands and French Polynesia (Table I) with A. Amansia tends to bleach to translucent white. an additional field test to distinguish Amansia spp. thicker. and trichoblasts lacking protective vesicles when young (Norris 1988b). On the other hand. Japan. while the other (US 04057b) is Melanamansia glomerata.

while younger plants have less cortication and more lax or absent rosettes of blades. alternate marginal proliferations (which represent non-determinate or endogenous secondary branchlets) 2–5 mm long (Fig. they were clearly apparent and about half the size of pericentral cells (Fig. more corticated secondary and primary axes with tight rosettes. M. These ventral. Indonesian and Indian Ocean region were sectioned and examined for pseudo-pericentral cells. Payri and N’Yeurt 1997) were reexamined. however it is clear that the degree of rosette formation and cortication of the axes is not a reliable taxonomic character at the species level in Amansia. when present). and found to consistently lack pseudo-pericentral cells and a similar observation was made on samples from Taiwan. an examination of the holotype and liquid-preserved and pressed isotypes of A. 1995). com. and found to lack pseudo-pericentral cells. Indian Ocean Amansia rhodantha also lacked such cells.). some specimens were virtually indistinguishable from specimens of A. paloloensis. the description of the species states that branches have the same shape as the main axis (i. glomerata occurs in the Solomon Islands.g. except the Hawaiian samples. and a main axis with alar cells not very distinct from secondary branches. fig. However. Fiji and New Caledonia (Table I). calm waters (such as blue holes. which was the main reason for separating the species from A.g. The illustration by Harvey of the holotype of Delesseria rhodantha (1834. pl. (2000: 188. both on the same plant and within individuals in similar populations. and in some instances lacking altogether. in some cases. Generally. Ecological studies would be needed to verify these hypotheses. and some samples from Kenya. fig. which also had a consistently thicker thallus. while in Malaysia Masuda et al. pseudo-pericentral cells were universally absent. degree of marginal serrations and thallus flattening. Wilson and Kraft 2000). 1) shows both the presence of tight and lax rosettes of secondary branches. type of endogenous branching. From an examination of the specimens at hand. the presence or absence of bladelets in rosettes. M. 19). in New Caledonia and Kenya). which is a characteristic feature of both Amansia and Melanamansia (Norris 1988b) can vary from lax to tight.Revision of some Amansia and Melanamansia species 239 recording of the Wilkes Expedition collections was not very satisfactory. and would seem to function as attachment haptera. 41) similarly report the absence of pseudo-pericentral cells in A. 7). Melanamansia glomerata is unlikely to be confused with other Melanamansia species because of its unique combination of characteristics (Norris 1988b. pers. more exposed conditions (such as on fringing reefs) favour tight rosette formation (which is more hydrodynamically resistant) while more lax arrangements are possible in deeper. thus placing them in Amansia. Fijian and French Polynesian specimens previously ascribed to Melanamansia (N’Yeurt et al. Masuda. rhodantha. L 0108603). In most instances. There is a possibility it might have been mixed with collections from neighbouring localities such as the Solomon Islands. the Philippines. Herbarium specimens of ‘Amansia glomerata’ from southern Japan housed in SAP (Hokkaido University) were examined. Proliferations from the ventral midrib (aside from secondary branchlets) were only found to be abundant in some of the Indonesian Amansia rhodantha specimens from Salayer (e. and the presence of tight and lax rosettes of secondary branches on the same plant (Figs 11–13). The presence or absence of these pseudo-pericentral cells hence appears to be a good generic criterion. both have an ala) and are arranged in ‘lax’ rosettes (South and Skelton 1999: 247. Indeed. environmental factors seem to play a role in determining the degree of rosette formation. older plants tend to have thicker. rhodantha from Mauritius and Western Australia (compare Figs 4. Also. and is backed by other morphological and biochemical differences (Norris 1988b. Herbarium and liquid-preserved samples of Amansia and Melanamansia species from the Pacific. 13). fig. perhaps an adaptation to the exposed habitats where the plants came from. It would seem that perhaps shallow. so it cannot be confirmed at this stage if M. position of the reproductive axes and the distinctness of the axial ring in rosette axes (Norris 1988a.e. in the case of Amansia paloloensis). adding to the confusion between the two genera. Norris (1988b: 211) reported that rosette axes in . 1996. In Melanamansia glomerata these marginal serrations were much less pronounced (but more regular than in Amansia rhodantha. All Amansia rhodantha specimens examined had fairly prominent. 5. in the case of A. irregularly dichotomous proliferations lack alar cells. When pseudo-pericentral cells were present. For instance. paloloensis revealed a wide range in the degree of rosette formations and cortication of the secondary axes and presence of alae on primary axes. However. glomerata occurs in the same habitat as Amansia rhodantha (e. the vesiculate (in Amansia) or non-vesiculate (in Melanamansia) nature of trichoblasts. Validity of current criteria to distinguish the Amansieae Characters that have proved useful in distinguishing species in the Amansieae include the presence or absence of pseudo-pericentral cells. it was observed that rosettes of blades along the secondary axes. CXXVI. rhodantha. 2). The author has unsuccessfully tried to obtain material from the latter locality. with the blades smooth and superficially reminiscent of those of some Sargassum species (Figs 22–24). and hence the precise locality and date of this Melanamansia collection from Fiji is unknown.

Cells of the blade alae in both Amansia and Melanamansia are regularly elongate-hexagonal. There appears to be no notable difference in the range of thallus size. paloloensis specimens studied. rhodantha studied. it was noted that in Melanamansia the range was more restricted. . An additional potentially useful character is the number of segments between lateral ‘veins’ on the blades. with the length of segments ranging from 80 to 120 µm. One feature which was noted in Melanamansia glomerata is that individual tetrasporangial stichidia are often laterally branched once or twice (Fig. 25. which was also recorded by Norris (1988b) for Hawaiian material.). with most specimens having segment lengths of about 100 µm. cortication of axes and rosette formation between specimens of Amansia rhodantha studied.3. the number ranged from 3 to 7. Fig. B. However. D. rhodantha specimens observed in this study. a number of preliminary conclusions can be made: A. but can also occur apically. At this stage there seems to be no largely significant difference in the number of segments between alternating veins in the specimens of A. only being more or less surrounded by cortication. usually 3 or 4 with an average of 3. Hence the distinctness of the axial ring was not found a reliable taxonomic character among the species examined. In A. as well as between A. N’Yeurt Amansia from Natal have an ill-defined axial row of cells. R. Norris (1995: 66) counted the number of segment cells between these alternating veins for several species of Melanamansia and Amansia. paloloensis. com.240 A. and occur in two overlapping and slightly offset layers radiating on either side of the midrib. as the axial ring was still discernible in all sections made. An examination of these elegant cells in species of Amansia (Figs 10. in that they are typically rhodomelaceous with pairs of tetrasporangia in up to ten rows. as well as more corticated axes. which are in fact determinate branches covered by the pericentral cell derivatives. 15). showing the currently known distribution. but the range is reportedly less than in Amansia rhodantha. 20). Tetrasporangial stichidia are basically similar in both Amansia rhodantha and Melanamansia glomerata. in elongate flattened and pinnate stichidia which are usually on lateral marginal branchlets. smaller and narrower blades. These differences reinforce the conspecificity of the Amansia rhodantha and A. Biogeographical map for Amansia rhodantha (circle) and Melanamansia glomerata (triangle).4 (Norris 1995). as well as the generic distinctness of Melanamansia glomerata. which was not seen in Amansia rhodantha (Figs 6. This also varied according to the degree of rosette formation and age of the plant. 16) and Melanamansia show no particular difference. Cover cells usually totally cover the developing sporangia. This feature was examined on a range of specimens of Amansia from the South Pacific and the Indian Ocean but no significant difference was seen in these specimens. rhodantha and A. blade dimensions. and found them relatively constant. These can be easily detected in the unrolled apical portions of the blades by the developing trichoblast terminating each such alternating ‘vein’ (Norris 1995 and pers. From Table I. In Melanamansia it averages 4. Melanamansia glomerata from all localities studied have consistently smoother.

J. 1981. J. Masuda pers. M.A. Robin South is also thanked for his helpful comments on an earlier draft of this manuscript. U. 2000. and J. with descriptions of some new and little known species. S. B. Scheltema & Holkema. Jaasund. Schofield. Rhodophyta.Revision of some Amansia and Melanamansia species 241 Conclusions The results of this study and those of South and Skelton (1999) and of Masuda et al. composed of up to 4 cells: Melanamansia glomerata. M. flora and vegetation. and possibly other localities which are not yet adequately investigated. V. Professors Michio Masuda and Tadao Yoshida (SAP – Hokkaido University. A. G. Pub. E. Handbook No 2. trichoblasts non-vesiculate. Phaeophyceae. J. 7th edition. Sci. 2000. Lamouroux.. 1834. 29: 1–38. Sci. Index Herbariorum. Rhodophyceae. K. Volume II. – Absence of paired dorsal pericentral cells. 24 pls. P. Falkenberg. sparse. Kenya (this study). Annal. Berlin. 1987. Decaisne. 1999. 22: 161–193. C. Coppejans. Marine botany of the Kenya coast. M. 1901. Bishop Museum Press. Lunda (Lund). J. Bohn. 458 figs.). M. Revised Key to the Species of Amansia and Melanamansia in the South Pacific – Presence of paired dorsal pericentral cells. Cal. xv + 673 pp. I. A. University of Tromsø. Bull. Gleerups Förlag. Bot. Honolulu. W. Lewis. Dublin).. Foreword. (2000) confirm that the genus Amansia does exist in the Pacific Ocean. Acknowledgements The author sincerely thanks the following persons and institutions for the generous loan of type material and specimens used in this study: Dr John Huisman (MURU – Murdoch University). Bot. J. 2. pigments reddish-brown. 1976. Mr Barrett Brooks and Dr John Sims (US – Smithsonian Institution. Nouv. 1956. J. Professor G. Kylin. K. Dr Claude Payri (UPF – Tahiti). Part 1. Ms Susanna Riebe (LD – Botanical Museum. xiv + 939 pp. 1988. 27: 7–28. 1956. Intertidal Seaweeds in Tanzania. Washington). Mar. It is clear that both A. and A. Dr R. C. S. New Caledonia and the Philippines (this study). Isaac. Bot. African Bot. Norris is thanked for his helpful insights and suggestions for this research. F. 173 + (2) pp. H. Utrecht. Bot. First edition. Die Rhodomelaceen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Shimada. Dr John Parnell (TCD – Trinity College. Robin South and Mr Posa A. E. Notice of a collection of algae. Skelton (SUVA – The University of the South Pacific). The Structure and Reproduction of the Algae. The Herbaria of the World. Marine Red Algae of the Hawaiian Islands. from “Cap Malheureux”. Kato. F. 10 figs. Bot. Hommersand. 1841. J. 1945. Lord Howe Island (G. Fritsch.. Isaac. T. 2 folded maps. 35: 165–366. Sapporo). E. Nat. xvi + 477 + [1] pp. com. Seven species of Rhodophyceae. (Hooker) 1: 147–157. Sci. K. pig- References Abbott. 1809. Harvey. Tsuda. Kawaguchi and S. initially vesiculate and composed of up to 8 cells: Amansia rhodantha. E. and each genus is consistently differentiated from the other by a number of anatomical and biochemical characters. Isaac. C. 43: 315–346. Keuken and E. pls v–vii. W. N. Marine red algae from the North coast of Papua New Guinea. Professor Willem Prud’homme van Reine (L – Rijksherbarium. Australian Coral Reef Society: Brisbane. Charles Telfair. xxxviii + 312 pp. Millar for his generous assistance in obtaining copies of rare publications. Observations sur la physiologie des algues marines. 1983. Marine Algae of the Southern Great Barrier Reef. rhodantha and Melanamansia glomerata are more widely distributed than previously thought. Leiden). K. 336 figs.A. Lund. Professor G. frontispiece. Norris. E. Garrigue. K. xvi + 754 pp. Hist. Mar. J. W.). com. Nat. The morphology and classification of some Ceramiaceae and Rhodomelaceae. Hooker by the late Mrs. cxxvi. 1968. and it appears that Melanamansia glomerata is so far restricted to Hawaii (Norris 1995). W. trichoblasts on every second or third axial cell. E. J. Mar. Holmgren. P. Linnaea 15: 1–50. Fauna und Flora des Golfes von Neapel. 17: 297–380. E. East Africa Nat. Contrib. in the Mauritius. Marine algae of Inhaca Island and of the Inhaca Peninsula. Micronesica 21: 53–70. 3. S. Smiths. et description de cinq nouveaux genres de cette famille. 1842. South pers. Taxonomic notes on marine algae from Malaysia. H. A history and annotated account of the benthic marine algae of Taiwan. II. 443–457. pls cxxv. Monographie 26. ser. and Dr A. Phang.. and R. 43: 181–190. Masuda. R.W. Catalogue of marine benthic algae from New Caledonia. I. Millar. Myxophyceae.Vietnam and Malaysia (M. and F. Part I. Mus. Soc. Accepted 17 January 2002.. Phil.. Agardh. Essais sur une classification des algues et des polypiers calcifères de Lamouroux. Die Gattungen der Rhodophyceen. Cambridge University Press. 1963. Cribb. In historiam algarum symbolae.. Agardh. Soc. 1824. Lund). ments rose-red. . Figure 25 presents a world distribution of these two genera as it is currently known. Systema algarum. H. Paris 1: 330–333. communicated to Dr. iii + 159 pp. General account of the environment. 71 pls.

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